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AFROTROPICAL 17th edition (2018). MARK C. WILLIAMS. http://www.lepsocafrica.org/?p=publications&s=atb

Genus Boisduval, 1832

Voyage de Découvertes de L’Astrolabe sous le commandement de M. J. dumont D’Urville. Faune entomologique de l’Océan Pacifique. Lépidopteres (1): 117 (267 pp.). Paris. Type-species: Papilio thyonneus Cramer, by subsequent designation (Scudder, 1875. Proceedings of the American Academy of Arts and Sciences 10: 152 (91-293).) [extralimital].

Synonyms based on extralimital type-species: Apsithra Moore; Sykophages Martin.

Cyrestis camillus camillus. Image courtesy of Jeremy Dobson.

The genus Cyrestis belongs to the Family Rafinesque, 1815; Subfamily Guenée, 1865. There are no other genera in the Subfamily Cyrestinae in the Afrotropical Region.

Cyrestis (Porcelains) is an Old World genus of 22 species, only one of which is Afrotropical.

Subgenus Azania Martin, 1903

Deutsche Entomologische Zeitschrift, Iris 16: 160 (71-169). Type-species: Papilio camillus Fabricius, by subsequent designation (Hemming, 1939. Proceedings of the Royal Entomological Society of London (B) 8: 133 (133-138).).

*Cyrestis (Azania) camillus (Fabricius, 1781)# African Porcelain

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Mudpuddling African butterflies ( ) in Tanzania (left) and Madagascar (right). Images courtesy Sudheer Kommana (left) and Bart Wursten (right).

Papilio camillus Fabricius, 1781. Species Insectorum 2: 11 (499 pp.). Hamburgi & Kilonii. Cyrestis (Azania) pantheus (Drury, 1782). Pringle et al., 1994: 111. [synonym of camillus]

Cyrestis camillus camillus. Male (Wingspan 47 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 15 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Cyrestis camillus camillus. Female (Wingspan 53 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 1 December 2007. J. Dobson. Images M.C. Williams ex Dobson Collection.

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Cyrestis camillus camillus. Male (melanic form). Left – upperside; right – underside. Central African Republic. March, 2014. Images M.C. Williams ex J. Lawrence Collection.

Alternative common name: African Map . Type locality: “Africa aequinoctiali”. Diagnosis: Some seasonal variation is shown by this otherwise very distinctive butterfly: the golden transverse bands on the underside of the hindwing are strongly marked in midsummer specimens but may be very faint in the early spring brood (Pringle et al., 1994). Distribution: Senegal, Guinea-Bissau (Bivar-de-Sousa et al., 2008), Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Nigeria, Cameroon, Equatorial Guinea, Sao Tome & Principe, Gabon, Congo, Angola, Central African Republic, Democratic Republic of Congo, Ethiopia, Uganda, Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, South Africa, Madagascar. Habitat: Forest and dense secondary growth. Commoner in secondary than in primary forest (Larsen, 2005a). In Tanzania the nominate subspecies occurs from 800 to 1 500 m and subspecies sublineata from near sea level to 1 600 m (Kielland, 1990d). Habits: Despite its small body, relative to the wing surface, this butterfly is a powerful flier. When not threatened the flight is often gliding, with occasional flaps of the wings, much as in the genus Neptis. Specimens are usually found in the canopy of trees near streams but males often come down to suck moisture at damp spots on the ground. When mudpuddling they assume a unique position, holding the wings open and inclined slightly downwards, with the forewings pushed forwards (Van Son, 1979). Males also come to foul substances and urine, running about excitedly with the wings held flat (Larsen, 2005a). If disturbed they will fly off rapidly, with a zig-zagging flight, and alight suddenly on the underside of a leaf with the wings held flat, thus appearing almost to miraculously disappear (Van Son, 1979). Individuals also alight with open wings on the upper surface of the leaves of branches that overhang water (Kielland, 1990d). Specimens are strongly attracted to flowers (Pringle et al., 1994). Van Son (1979) states that when settled it resembles the distasteful uraniid moth Urapteroides falcifera Warren. R. Schutte (pers. comm. April 2010) found ssp. elegans in Madagascar to be a common resident, mud puddling with swarms of pierids along a forest track. Its open wing gliding flight made it easily distinguishable from the other white butterflies in the swarm. Flight period: All year, with slight seasonal variation. Early stages:

Mullin, in Pringle et al., 1994: 111 [as Cyrestis pantheus sublineatus; Mount Selinda, Zimbabwe; first bred in February, 1989]. “Egg: Eggs are generally laid on the edges of young leaves or on new buds. The egg is bright yellow when laid. It measures 0.7 mm wide at the base by 0.9 mm high. It is elliptical in shape with a flattened and recessed top. It has 9-11 longitudinal ribs. After a couple of days the egg develops brownish streaks between the ribs, giving it an orange appearance. The eggs hatched after some 10 days (much slower than the summer brood). Larva: The first instar larva is 1.5 mm long, a watery greenish-ochre colour with a black head. It devours its eggshell for the first meal. The anal segment is wine-brown and bears a short sphingid-like horn. Final instar larva: Approximately 40 mm long (maximum size). Head: spherical, bearing a pair of horns which curve outwards and backward, similar to a

3 young goat, about 5 mm long; face cream edged in grape-black; horns grape-black, and this coloration extends in two diagonal stripes to link with the mouth parts. The body exhibits a tan-coloured dorsal strip extending from head to tail. Segments 1, 2 and 3 dorsally tan-coloured merging to plum-black laterally. From the fourth segment arises a backward-pointing curved horn, about 8 mm long, plum-black in colour as is the dorsum to the tail. Lateral coloration bright, pale green to the tail with the exception of segments 5 and 7, which are grape-black with a black diagonal stripe extending dorsally backwards into segments 6 and 8. A very narrow, faint diagonal stripe may also be discerned on segments 4, 6, 8, 9 and 10. Arising from the anal segment is a backward-pointing horn which then curves forward before curving yet again backwards at the tip rather like an S-bend. The final instar larvae rest on the upper surface of the leaves, relying no doubt on their coloration as camouflage against predators. Pupa: Medium brown; 28 mm long; similar to a Neptis; palps protrude at the anterior end, and these are not joined although they meet; slightly winged laterally. The thoracic area is dorsally domed and pointed and embellished, where it joins the abdomen, with a silver spot on either side of the dorsal line. A dried leaf-like structure appears dorsally behind the thorax in a vertical plane, and the remaining abdominal segments are serrated dorsally, and to a lesser extent, ventrally. The anal segment is laterally flattened and dark brown. The whole pupa is netted with veining which resembles a semi-decomposed leaf. Although individual durations were not recorded, the whole cycle from egg- laying to adult emergence took approximately five weeks. Young larvae seem to cut main leaf veins to the area on which they are feeding, probably to prevent the sap flow. First instar larvae make ‘ropes’ in the fashion of Pseudacraea boisduvalii trimenii. The ropes are black and the larva rests at the end”.

Larval food: Antiaris toxicaria Lesch. var. africana Scott-Elliot ex A.Chev. (Moraceae) [Vuattoux & Blandin, 1979; as Antiaris africana; Ivory Coast]. madagascariense DC. (Moraceae) [Mullin, in Pringle et al., 1994: 112; as Bosquiea phoberos Baill.; Mount Selinda, Zimbabwe]. Milicia excelsa (Welw.) C.C.Berg. (Moraceae) [Vuattoux & Blandin, 1979; as Chlorophora excelsa; Ivory Coast]. Chlorophora species (Moraceae) [Owen, vide Larsen, 2005; Sierra Leone]. sur Forssk. (Moraceae) [Vuattoux & Blandin, 1979; as Ficus capensis; Ivory Coast]. Ficus exasperata Vahl (Moraceae) [Vuattoux & Blandin, 1979; Ivory Coast]. Forssk. (Moraceae) [Mullin, in Pringle et al., 1994: 112; Chimanimani, Zimbabwe]. mesozygia Stapf (Moraceae) [Vuattoux & Blandin, 1979; Ivory Coast]. Treculia species (Moraceae) [Pyrcz, 1991; Principe]. Trilepisium madagascariense DC. (Moraceae) [Mullin, in Pringle et al., 1994: 112]. species (Rhamnaceae) [S. Collins, vide Larsen, 1991c; Kenya coast].

Note: Cyrestis pantheus pantheus (Drury, 1782) was proposed as replacement name for Papilio camillus Fabricius, 1781, nec Cramer, 1780 – see Koçak, 1996 (Centre for Entomological Studies Miscellaneous Papers 27-28: 12 (10- 16). Proposed conservation of Cyrestis camillus by Larsen, 1997 was upheld (Opinion 1917, ICZN 1999) – see Larsen, 1997 (Bulletin of Zoological Nomenclature 54 (3): 157 (155-158)).

Cyrestis (Azania) camillus camillus (Fabricius, 1781)

Papilio camillus Fabricius, 1781. Species Insectorum 2: 11 (499 pp.). Hamburgi & Kilonii.

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Cyrestis camillus camillus. Male (Wingspan 47 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 15 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Cyrestis camillus camillus. Female (Wingspan 53 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 1 December 2007. J. Dobson. Images M.C. Williams ex Dobson Collection.

Cyrestis camillus camillus. Male (melanic form). Left – upperside; right – underside. Central African Republic. March, 2014. Images M.C. Williams ex J. Lawrence Collection.

Type locality: “Africa aequinoctiali”. Distribution: Senegal, Guinea-Bissau (Bivar-de-Sousa et al., 2008), Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Nigeria (south and Cross River loop), Cameroon, Equatorial Guinea (Bioko), Sao Tome & Principe (Principe), Gabon, Congo, Angola, Central African Republic, Democratic Republic of Congo, Ethiopia, Uganda, Kenya (west), Tanzania (north-west), Zambia (north). 5 Specific localities: Senegal – Basse Casamance (single record) (Larsen, 2005a). Ghana – Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Cameroon – Korup (Larsen, 2005a). Equatorial Guinea – Caldera de Luba, Bioko (Martin, 2015). Gabon – Lake Evaro (Vande weghe, 2010); Kinguele (Vande weghe, 2010); Waka (Vande weghe, 2010); Bitam (Vande weghe, 2010); Ipassa (Vande weghe, 2010); Franceville (Vande weghe, 2010); Ekouyi (Vande weghe, 2010); camp PPG (Vande weghe, 2010). Central African Republic – Dzanga (Noss, 1998). Democratic Republic of Congo – Stanleyville (Le Cerf, 1927). Uganda – Semuliki N.P. (Davenport & Howard, 1996). Tanzania – Mpanda and Kigoma Districts (Kielland, 1990d).

pantheus Drury, 1782 (as sp. of Papilio). Illustrations of Natural History 3: index et 7 (76 pp.). London. Sierra Leone: “Sierra Leon”.

donckieri Le Cerf, 1927 (as f. (? indiv.) of Cyrestis camillus). Encyclopédie Entomologique (B. 3. ) 2: 48 (44-58). [Democratic Republic of Congo]: “Stanleyville, Congo belge”.

Cyrestis (Azania) camillus elegans Boisduval, 1833

Cyrestis elegans Boisduval, 1833. Nouvelles Annales du Muséum d’Histoire Naturelle, Paris 2: 190 (149-270).

Cyrestis camillus elegans. Male. Left – upperside; right – underside. Madagascar. May, 2015. Images M.C. Williams ex J. Lawrence Collection.

Type locality: Madagascar: “à Sainte-Marie, à Foule-Pointe, et à Tamatave [Toamasina]”. Distribution: Madagascar. Specific localities: Madagascar – Sainte-Marie (TL); Foule-Pointe (Boisduval, 1833), Toamasina (Boisduval, 1833); Perinet Montadia primary forest (R. Schutte, pers. comm. April 2010).

Cyrestis (Azania) camillus sublineata Lathy, 1901#

6 Cyrestis sublineata Lathy, 1901. Transactions of the Entomological Society of London 1901: 25 (19-36). Cyrestis camillus sublineatus Lathy, 1901. Dickson & Kroon, 1978. Cyrestis (Azania) pantheus sublineatus Lathy, 1901. Pringle et al., 1994: 111.

Cyrestis camillus sublineata. Male. Left – upperside; right – underside. Essex Valley, Vumba, Zimbabwe. 17 December 1998. J. Greyling. Images M.C. Williams ex Greyling Collection.

Cyrestis camillus sublineata. Female. Left – upperside; right – underside. Rusito, Zimbabwe. 22 September 1996. J. Greyling. Images M.C. Williams ex Greyling Collection.

Type locality: [Malawi]: “Zomba”. Diagnosis: Differs from the nominate subspecies in that the forewing orange discal band is irregular (in the nominate subspecies it is straight to vein 4) (Kielland, 1990d). Distribution: Kenya (east), Tanzania (east), Malawi, Mozambique, Zimbabwe (eastern border), South Africa (Limpopo Province). Recorded, apparently in error, from Zambia by Ackery et al., 1995 (Heath et al., 2002). Specific localities: Kenya – coast (Larsen, 1991c); Teita Hills (Larsen, 1991c); Shimba Hills (Larsen, 1991c). Tanzania – Rondo (Kielland, 1990d); Tanga Region (Kielland, 1990d); Pare Mountains (Kielland, 1990d); Mount Meru (Kielland, 1990d); Moshi (Kielland, 1990d); Morogoro (Kielland, 1990d); Uzungwa Range (Kielland, 1990d); Masagati Forest (Kielland, 1990d). Malawi – Zomba (TL); Mt Mulanje (Congdon et al., 2010). Mozambique – Dondo (Pringle et al., 1994); Xiluvo (Pringle et al., 1994); Maronga (Pringle et al., 1994); Posto de Catandica (Pringle et al., 1994); Mt Mabu (Congdon et al., 2010). Zimbabwe – Mutare (Pringle et al., 1994); Mount Selinda (Pringle et al., 1994); Shamvu, north-east of

7 Harare (Paré); junction of Sabi and Runde rivers (Pringle et al., 1994); Chiredzi River (Mullin). Limpopo Province – Pafuri (Pringle et al., 1994); Punda Maria (Kloppers); Woodbush (Swanepoel, 1983; single record); Lekgalameetse Nature Reserve (“Malta Forest”) (Badham, vide Swanepoel (1983); single record).

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